Predicting the distribution of four species of raptors (Aves: Accipitridae) in southern Spain: statistical models work better than existing maps

Aim To test the effectiveness of statistical models based on explanatory environmental variables vs. existing distribution information (maps and breeding atlas), for predicting the distribution of four species of raptors (family Accipitridae): common buzzard Buteo buteo (Linnaeus, 1758), short‐toed eagle Circaetus gallicus (Gmelin, 1788), booted eagle Hieraaetus pennatus (Gmelin, 1788) and black kite Milvus migrans (Boddaert, 1783). Location Andalusia, southern Spain. Methods Generalized linear models of 10 × 10 km squares surveyed for the presence/absence of the species by road census. Statistical models use as predictors variables derived from topography, vegetation and land‐use, and the geographical coordinates (to take account of possible spatial trends). Predictions from the models are compared with current distribution maps from the national breeding atlas and leading reference works. Results The maps derived from statistical models for all four species were more predictive than the previously published range maps and the recent national breeding atlas. The best models incorporated both topographic and vegetation and land‐use variables. Further, in three of the four species the inclusion of spatial coordinates to account for neighbourhood effects improved these models. Models for the common buzzard and black kite were highly predictive and easy to interpret from an ecological point of view, while models for short‐toed eagle and, particularly, booted eagle were not so easy to interpret, but still predicted better than previous distribution information. Main conclusions It is possible to build accurate predictive models for raptor distribution with a limited field survey using as predictors environmental variables derived from digital maps. These models integrated in a geographical information system produced distribution maps that were more accurate than previously published ones for the study species in the study area. Our study is an example of a methodology that could be used for many taxa and areas to improve unreliable distribution information.     

Spatial variation in the climatic predictors of species compositional turnover and endemism

Previous research focusing on broad‐scale or geographically invariant species‐environment dependencies suggest that temperature‐related variables explain more of the variation in reptile distributions than precipitation. However, species–environment relationships may exhibit considerable spatial variation contingent upon the geographic nuances that vary between locations. Broad‐scale, geographically invariant analyses may mask this local variation and their findings may not generalize to different locations at local scales. We assess how reptile–climatic relationships change with varying spatial scale, location, and direction. Since the spatial distributions of diversity and endemism hotspots differ for other species groups, we also assess whether reptile species turnover and endemism hotspots are influenced differently by climatic predictors. Using New Zealand reptiles as an example, the variation in species turnover, endemism and turnover in climatic variables was measured using directional moving window analyses, rotated through 360°. Correlations between the species turnover, endemism and climatic turnover results generated by each rotation of the moving window were analysed using multivariate generalized linear models applied at national, regional, and local scales. At national‐scale, temperature turnover consistently exhibited the greatest influence on species turnover and endemism, but model predictive capacity was low (typically r² = 0.05, P < 0.001). At regional scales the relative influence of temperature and precipitation turnover varied between regions, although model predictive capacity was also generally low. Climatic turnover was considerably more predictive of species turnover and endemism at local scales (e.g., r² = 0.65, P < 0.001). While temperature turnover had the greatest effect in one locale (the northern North Island), there was substantial variation in the relative influence of temperature and precipitation predictors in the remaining four locales. Species turnover and endemism hotspots often occurred in different locations. Climatic predictors had a smaller influence on endemism. Our results caution against assuming that variability in temperature will always be most predictive of reptile biodiversity across different spatial scales, locations and directions. The influence of climatic turnover on the species turnover and endemism of other taxa may exhibit similar patterns of spatial variation. Such intricate variation might be discerned more readily if studies at broad scales are complemented by geographically variant, local‐scale analyses.     

From topography to hydrology—The modifiable area unit problem impacts freshwater species distribution models

Species distribution models (SDMs) are statistical tools to identify potentially suitable habitats for species. For SDMs in river ecosystems, species occurrences and predictor data are often aggregated across subcatchments that serve as modeling units. The level of aggregation (i.e., model resolution) influences the statistical relationships between species occurrences and environmental predictors—a phenomenon known as the modifiable area unit problem (MAUP), making model outputs directly contingent on the model resolution. Here, we test how model performance, predictor importance, and the spatial congruence of species predictions depend on the model resolution (i.e., average subcatchment size) of SDMs. We modeled the potential habitat suitability of 50 native fish species in the upper Danube catchment at 10 different model resolutions. Model resolutions were derived using a 90‐m digital‐elevation model by using the GRASS‐GIS module r.watershed. Here, we decreased the average subcatchment size gradually from 632 to 2 km². We then ran ensemble SDMs based on five algorithms using topographical, climatic, hydrological, and land‐use predictors for each species and resolution. Model evaluation scores were consistently high, as sensitivity and True Skill Statistic values ranged from 86.1–93.2 and 0.61–0.73, respectively. The most contributing predictor changed from topography at coarse, to hydrology at fine resolutions. Climate predictors played an intermediate role for all resolutions, while land use was of little importance. Regarding the predicted habitat suitability, we identified a spatial filtering from coarse to intermediate resolutions. The predicted habitat suitability within a coarse resolution was not ported to all smaller, nested subcatchments, but only to a fraction that held the suitable environmental conditions. Across finer resolutions, the mapped predictions were spatially congruent without such filter effect. We show that freshwater SDM predictions can have consistently high evaluation scores while mapped predictions differ significantly and are highly contingent on the underlying subcatchment size. We encourage building freshwater SDMs across multiple catchment sizes, to assess model variability and uncertainties in model outcomes emerging from the MAUP.     

Tree species distribution in temperate forests is more influenced by soil than by climate

Knowledge of the ecological requirements determining tree species distributions is a precondition for sustainable forest management. At present, the abiotic requirements and the relative importance of the different abiotic factors are still unclear for many temperate tree species. We therefore investigated the relative importance of climatic and edaphic factors for the abundance of 12 temperate tree species along environmental gradients. Our investigations are based on data from 1,075 forest stands across Switzerland including the cold‐induced tree line of all studied species and the drought‐induced range boundaries of several species. Four climatic and four edaphic predictors represented the important growth factors temperature, water supply, nutrient availability, and soil aeration. The climatic predictors were derived from the meteorological network of MeteoSwiss, and the edaphic predictors were available from soil profiles. Species cover abundances were recorded in field surveys. The explanatory power of the predictors was assessed by variation partitioning analyses with generalized linear models. For six of the 12 species, edaphic predictors were more important than climatic predictors in shaping species distribution. Over all species, abundances depended mainly on nutrient availability, followed by temperature, water supply, and soil aeration. The often co‐occurring species responded similar to these growth factors. Drought turned out to be a determinant of the lower range boundary for some species. We conclude that over all 12 studied tree species, soil properties were more important than climate variables in shaping tree species distribution. The inclusion of appropriate soil variables in species distribution models allowed to better explain species' ecological niches. Moreover, our study revealed that the ecological requirements of tree species assessed in local field studies and in experiments are valid at larger scales across Switzerland.     

Estimation of climatic favourable areas for locust outbreaks in Spain: integrating species' presence records and spatial information on outbreaks

For millennia, locust swarms have recurrently devastated crop productivity across continents. In much of Europe, locust outbreaks have been considerably reduced by human pressure in recent decades, but important foci of outbreaks still exist in Spain. Distribution models are often used to derive spatial hypotheses and risk maps. Because insufficient information is available to include the extreme plasticity of the solitary and gregarious phases of locusts in large‐scale spatial models, we modelled the distribution either of Acrididae species or of outbreaks per se. Confirmed occurrences of Dociostaurus, Calliptamus and Chorthippus species were obtained from a field survey complemented by museum collection data and the published literature. The locations of confirmed or potential outbreaks covering two time periods of 20 years each were obtained from the literature and from Spanish autonomous community reports. Models were built with one topographic and eleven climatic predictors. We evaluated the ability of different models to predict outbreak recurrence and found that models based on Moroccan locust data or outbreak occurrence data performed the best. We generated a predictive map of the climatic favourability for locust outbreaks in Spain and found that the major foci of locust swarms were encompassed by those areas categorized by the models as areas of highest risk. Predictive maps of outbreak favourability can facilitate the more sustainable use of insecticides and more efficient integrated pest management.     

Spatial modelling of ecological indicator values improves predictions of plant distributions in complex landscapes

Ecologically meaningful predictors are often neglected in plant distribution studies, resulting in incomplete niche quantification and low predictive power of species distribution models (SDMs). Because environmental data are rare and expensive to collect, and because their relationship with local climatic and topographic conditions are complex, mapping them over large geographic extents and at high spatial resolution remains a major challenge. Here, we propose to derive environmental data layers by mapping ecological indicator values in space. We combined ~6 million plant occurrences with expert-based plant ecological indicator values (EIVs) of 3600 species in Switzerland. EIVs representing local soil properties (pH, moisture, moisture variability, aeration, humus and nutrients) and climatic conditions (continentality, light) were modelled at 93 m spatial resolution with the Random Forest algorithm and 16 predictors representing meso-climate, land use, topography and geology. Models were evaluated and predictions of EIVs were compared with soil inventory data. We mapped each EIV separately and evaluated EIV importance in explaining the distribution of 500 plant species using SDMs with a set of 30 environmental predictors. Finally, we tested how they improve an ensemble of SDMs compared to a standard set of predictors for ca 60 plant species. All EIV models showed excellent performance (|r| > 0.9) and predictions were correlated reasonably (|r| > 0.4) to soil properties measured in the field. Resulting EIV maps were among the most important predictors in SDMs. Also, in ensemble SDMs overall predictive performance increased, mainly through improved model specificity reducing species range overestimation. Combining large citizen science databases to expert-based EIVs is a powerful and cost–effective approach for generalizing local edaphic and climatic conditions over large areas. Producing ecologically meaningful predictors is a first step for generating better predictions of species distribution which is of main importance for decision makers in conservation and environmental management projects.     

Is current climatic equilibrium a guarantee for the transferability of distribution model predictions? A case study of the spotted hyena

Aim The transferability of species distribution models requires that species show climatic equilibrium throughout their entire distribution area. We test this assumption for the case of the spotted hyena, Crocuta crocuta, a large carnivore that has shifted its distribution over the last 100,000 years from a widespread Eurasian and African range to its current geographical distribution, restricted to the Sub‐Saharan areas of the African continent. Location Western Eurasia and Africa. Methods The current realized distribution of C. crocuta was estimated using presences and reliable absences as well as climatic, land‐cover and anthropic variables as predictors. The potential distribution was estimated using presences and a set of pseudo‐absences selected from localities outside climatically suitable localities, with only climatic variables serving as predictors. The current potential distribution was transferred to the Last Interglacial period (126,000 yr bp ) using the palaeoclimatic data yielded by the GENESIS 2 general circulation model, and validated with European fossil data. Generalized linear models were used on all occasions. Results Climatic variables are able to predict the current distribution of the species with high accuracy. The geographical projection of this model indicates that the species is distributed over almost all of its potential suitable area, which allows us to suppose that the current distribution of this species is in climatic equilibrium. However, the time transference of model predictions for the western Eurasian region reveals almost no suitable conditions for hyenas, despite the widespread presence of C. crocuta fossil remains on this continent during the Last Interglacial period. Main conclusions Our results indicate that, even when model results suggest a climatic equilibrium for a species distribution, the time transferability of such models does not necessarily provide realistic results. This occurs because the current geographical range does not allow estimations of all of the environmental requirements of a species. Therefore, any model trained with current data risks underestimating the potential suitable environmental and geographical range for species in a new area or time period.     

Geographic range estimates and environmental requirements for the harpy eagle derived from spatial models of current and past distribution

Understanding species–environment relationships is key to defining the spatial structure of species distributions and develop effective conservation plans. However, for many species, this baseline information does not exist. With reliable presence data, spatial models that predict geographic ranges and identify environmental processes regulating distribution are a cost‐effective and rapid method to achieve this. Yet these spatial models are lacking for many rare and threatened species, particularly in tropical regions. The harpy eagle (Harpia harpyja) is a Neotropical forest raptor of conservation concern with a continental distribution across lowland tropical forests in Central and South America. Currently, the harpy eagle faces threats from habitat loss and persecution and is categorized as Near‐Threatened by the International Union for the Conservation of Nature (IUCN). Within a point process modeling (PPM) framework, we use presence‐only occurrences with climatic and topographical predictors to estimate current and past distributions and define environmental requirements using Ecological Niche Factor Analysis. The current PPM prediction had high calibration accuracy (Continuous Boyce Index = 0.838) and was robust to null expectations (pROC ratio = 1.407). Three predictors contributed 96% to the PPM prediction, with Climatic Moisture Index the most important (72.1%), followed by minimum temperature of the warmest month (15.6%) and Terrain Roughness Index (8.3%). Assessing distribution in environmental space confirmed the same predictors explaining distribution, along with precipitation in the wettest month. Our reclassified binary model estimated a current range size 11% smaller than the current IUCN range polygon. Paleoclimatic projections combined with the current model predicted stable climatic refugia in the central Amazon, Guyana, eastern Colombia, and Panama. We propose a data‐driven geographic range to complement the current IUCN range estimate and that despite its continental distribution, this tropical forest raptor is highly specialized to specific environmental requirements.     

Importance of soil nutrients in the distribution of forest communities on a large geographical scale

Aim Soil nutrient content plays a key role in plant growth through mineral nutrition and toxicity. Its impact on plant species and community distribution is studied on a large geographical scale through surrogates like topography or geology. We investigated the importance of soil pH and C:N ratio, as direct nutritional gradients, to determine, with climatic factors, the spatial distribution of plant communities over large territories. Location We studied the distribution of six beech habitats of the NATURA 2000 network throughout France (550,000 km²). Methods Models were calibrated with 2108 floristic plots classified in the NATURA 2000 system and including climatic and topographic variables and soil nutritional measurements carried out in a laboratory. Logistic regression was used to model habitat distribution according to environmental variables. Climatic layers, a digital elevation model and maps of soil pH and nitrogen content, created using plant indicator values and large floristic databases, were used to map the sites suitable for beech communities. Distribution models were evaluated with an independent set of 2091 phytosociological plots. Results pH and nitrogen supply were the key distribution drivers for four of the six beech communities on a national scale. Their use in the distribution models distinguished within homogeneous climatic territories a gradient of nutritional conditions from acidic areas, suitable for nutrient‐poor beech communities, to calcareous areas suitable for nutrient‐rich ones. Predicted maps of beech habitats fit the spatial distribution of validation plots. Main conclusions Soil pH and nitrogen supply strongly improve predictions of forest community distribution carried out with climatic variables on a broad geographical scale. They allow delineation of areas with nutritional conditions suitable for each community, as well as the realization of predictive high‐resolution maps over large areas useful for sustainable and conservation management. Nomenclature Tutin & Heywood (2001 ) Flora Europaea. Cambridge University Press, Cambridge.     

Habitat distribution models for intertidal seaweeds: responses to climatic and non‐climatic drivers

Aim Because intertidal organisms often live close to their physiological tolerance limits, they are potentially sensitive indicators of climate‐driven changes in the environment. The goals of this study were to assess the effect of climatic and non‐climatic factors on the geographical distribution of intertidal macroalgae, and to predict future distributions under different climate‐warming scenarios. Location North‐western Iberian Peninsula, southern Europe. Methods We developed distribution models for six ecologically important intertidal seaweed species. Occurrence and microhabitat data were sampled at 1‐km² resolution and analysed with climate variables measured at larger spatial scales. We used generalized linear models and applied the deviance and Bayesian information criterion to model the relationship between environmental variables and the distribution of each target species. We also used hierarchical partitioning (HP) to identify predictor variables with higher independent explanatory power. Results The distributions of Himanthalia elongata and Bifurcaria bifurcata were correlated with measures of terrestrial and marine climate, although in opposite directions. Model projections under two warming scenarios indicated the extinction of the former at a faster rate in the Cantabrian Sea (northern Spain) than in the Atlantic (west). In contrast, these models predicted an increase in the occurrence of B. bifurcata in both areas. The occurrences of Ascophyllum nodosum and Pelvetia canaliculata, species showing rather static historical distributions, were related to specific non‐climatic environmental conditions and locations, such as the location of sheltered sites. At the southernmost distributional limit, these habitats may present favourable microclimatic conditions or provide refuges from competitors or natural enemies. Model performances for Fucus vesiculosus and F. serratus were similar and poor, but several climatic variables influenced the occurrence of the latter in the HP analyses. Main conclusions The correlation between species distributions and climate was evident for two species, whereas the distributions of the others were associated with non‐climatic predictors. We hypothesize that the distribution of F. serratus responds to diverse combinations of factors in different sections of the north‐west Iberian Peninsula. Our study shows how the response of species distributions to climatic and non‐climatic variables may be complex and vary geographically. Our analyses also highlight the difficulty of making predictions based solely on variation in climatic factors measured at coarse spatial scales.     

Patterns of woody plant species richness in the Iberian Peninsula: environmental range and spatial scale

Aim Climate‐based models often explain most of the variation in species richness along broad‐scale geographical gradients. We aim to: (1) test predictions of woody plant species richness on a regional spatial extent deduced from macro‐scale models based on water–energy dynamics; (2) test if the length of the climate gradients will determine whether the relationship with woody species richness is monotonic or unimodal; and (3) evaluate the explanatory power of a previously proposed ‘water–energy’ model and regional models at two grain sizes. Location The Iberian Peninsula. Methods We estimated woody plant species richness on grid maps with c. 2500 and 22,500 km² cell size, using geocoded data for the individual species. Generalized additive models were used to explore the relationships between richness and climatic, topographical and substrate variables. Ordinary least squares regression was used to compare regional and more general water–energy models in relation to grain size. Variation partitioning by partial regression was applied to find how much of the variation in richness was related to spatial variables, explanatory variables and the overlap between these two. Results Water–energy dynamics generate important underlying gradients that determine the woody species richness even over a short spatial extent. The relationships between richness and the energy variables were linear to curvilinear, whereas those with precipitation were nonlinear and non‐monotonic. Only a small fraction of the spatially structured variation in woody species richness cannot be accounted for by the fitted variables related to climate, substrate and topography. The regional models accounted for higher variation in species richness than the water–energy models, although the water–energy model including topography performed well at the larger grain size. Elevation range was the most important predictor at all scales, probably because it corrects for ‘climatic error’ due to the unrealistic assumption that mean climate values are evenly distributed in the large grid cells. Minimum monthly potential evapotranspiration was the best climatic predictor at the larger grain size, but actual evapotranspiration was best at the smaller grain size. Energy variables were more important than precipitation individually. Precipitation was not a significant variable at the larger grain size when examined on its own, but was highly significant when an interaction term between itself and substrate was included in the model. Main conclusions The significance of range in elevation is probably because it corresponds to several aspects that may influence species diversity, such as climatic variability within grid cells, enhanced surface area, and location for refugia. The relative explanatory power of energy and water variables was high, and was influenced by the length of the climate gradient, substrate and grain size of the analysis. Energy appeared to have more influence than precipitation, but water availability is also determined by energy, substrate and topographic relief.     

Integrating dynamic environmental predictors and species occurrences: Toward true dynamic species distribution models

While biological distributions are not static and change/evolve through space and time, nonstationarity of climatic and land‐use conditions is frequently neglected in species distribution models. Even recent techniques accounting for spatiotemporal variation of species occurrence basically consider the environmental predictors as static; specifically, in most studies using species distribution models, predictor values are averaged over a 50‐ or 30‐year time period. This could lead to a strong bias due to monthly/annual variation between the climatic conditions in which species' locations were recorded and those used to develop species distribution models or even a complete mismatch if locations have been recorded more recently. Moreover, the impact of land‐use change has only recently begun to be fully explored in species distribution models, but again without considering year‐specific values. Excluding dynamic climate and land‐use predictors could provide misleading estimation of species distribution. In recent years, however, open‐access spatially explicit databases that provide high‐resolution monthly and annual variation in climate (for the period 1901–2016) and land‐use (for the period 1992–2015) conditions at a global scale have become available. Combining species locations collected in a given month of a given year with the relative climatic and land‐use predictors derived from these datasets would thus lead to the development of true dynamic species distribution models (D‐SDMs), improving predictive accuracy and avoiding mismatch between species locations and predictor variables. Thus, we strongly encourage modelers to develop D‐SDMs using month‐ and year‐specific climatic data as well as year‐specific land‐use data that match the period in which species data were collected.     

Quantitative prediction of the distribution and abundance of Vaccinium myrtillus with climatic and edaphic factors

Question: Can the distribution and abundance of Vaccinium myrtillus be reasonably predicted with soil nutritional and climatic factors? Location: Forests of France. Methods: We used Braun‐Blanquet abundance/dominance information for Vaccinium myrtillus on 2905 forest sites extracted from the phyto‐ecological database EcoPlant, to characterize the species ecological response to climatic and edaphic factors and to predict its cover/abundance at the national scale. The link between cover/abundance of the species and climatic (65 monthly and annual predictors concerning temperature, precipitation, radiation, potential evapotranspiration, water balance) and edaphic (two predictors: soil pH and C:N ratio) factors was investigated with proportional odds models. We evaluated the quality of our model with 9830 independent relevés extracted from Sophy, a large phytosociological database for France. Results: In France, Vaccinium myrtillus is at the southern limit of its European geographic range and three environmental factors (mean annual temperature, soil pH and C:N ratio) allow prediction of its distribution and abundance in forests with high success rates. The species reveals a preference for colder sites (especially mountains) and nutritionally poor soils (low pH and high C:N ratio). A predictive map of its geographic range reveals that the main potential habitats are mountains and northwestern France. The potential habitats with maximal expected abundance are the Vosges and the Massif central mountains, which are both acidic mountains. Conclusions: Complete niche models including climate and soil nutritional conditions allow an improvement of the spatial prediction of plant species abundance at a broad scale. The use of soil nutritional variables in distribution models further leads to an improvement in the prediction of plant species habitats within their geographical range.     

Response of female beetles to LIDAR derived topographic variables in Eastern boreal mixedwood forests (Coleoptera, Carabidae)

Biodiversity monitoring is increasingly being bolstered with high resolution data derived from remote sensing such as LIDAR (Light Detection and Ranging). We derived a series of topographical variables, including slope, azimuth, ground curvature and flow accumulation from LIDAR images and compared these to captures of female carabids in pitfall traps in Eastern boreal mixedwood forests. We developed a series of species-specific logistic models predicting the proportion of females for eight dominant species, including Agonum retractum, Calathus ingratus, Platynus decentis, Pterostichus adstrictus, Pterostichus coracinus, Pterostichus pensylvanicus, Sphaeroderus nitidicollis and Synuchus impunctatus. We used these models to test three hypotheses related to how the modest topography in boreal forests could influence the availability of microhabitats and possibly potential sites for oviposition and larval development. In general, topographic features such as north facing slopes and high flow accumulation were important predictors of the proportion of females. Models derived from larger scale topography, such as hillsides or small watersheds on the order of ¼-1 ha were better predictors of the proportion of females than were models derived from finer scale topography such as hummocks and small depressions. We conclude that topography likely influences the distribution of carabids based on hydrological mechanisms rather than factors related to temperature. We further suggest based on the scale of responses that these hydrological mechanisms may be linked to the attenuation of past disturbances by wildfire and the propensity of unburned forest patches and fire skips.     

Squamate richness in the Brazilian Cerrado and its environmental–climatic associations

We investigate patterns of species richness of squamates (lizards, snakes, and amphisbaenians) in the Brazilian Cerrado, identifying areas of particularly high richness, and testing predictions of large‐scale richness hypotheses by analysing the relationship between species richness and environmental climatic variables. We used point localities from museum collections to produce maps of the predicted distributions for 237 Cerrado squamate species, using niche‐modelling techniques. We superimposed distributions of all species on a composite map, depicting richness across the ecosystem. Then, we performed a multiple regression analysis using eigenvector‐based spatial filtering (Principal Coordinate of Neighbour Matrices) to assess environmental–climatic variables that are best predictors of species richness. We found that the environmental–climatic and spatial filters multiple regression model explained 78% of the variation in Cerrado squamate richness (r² = 0.78; F = 32.66; P < 0.01). Best predictors of species richness were: annual precipitation, precipitation seasonality, altitude, net primary productivity, and precipitation during the driest quarter. A model selection approach revealed that several mechanisms related to the different diversity hypothesis might work together to explain richness variation in the Cerrado. Areas of higher species richness in Cerrado were located mainly in the south‐west, north, extreme east, and scattered areas in the north‐west portions of the biome. Partitioning of energy among species, habitat differentiation, and tolerance to variable environments may be the primary ecological factors determining variation in squamate richness across the Cerrado. High richness areas in northern Cerrado, predicted by our models, are still poorly sampled, and biological surveys are warranted in that region. The south‐western region of the Cerrado exhibits high species richness and is also undergoing high levels of deforestation. Therefore, maintenance of existing reserves, establishment of ecological corridors among reserves, and creation of new reserves are urgently needed to ensure conservation of species in these areas.     

Climatic water deficit,tree species ranges,and climate change in Yosemite National Park

Aim (1) To calculate annual potential evapotranspiration (PET), actual evapotranspiration (AET) and climatic water deficit (Deficit) with high spatial resolution; (2) to describe distributions for 17 tree species over a 2300‐m elevation gradient in a 3000‐km² landscape relative to AET and Deficit; (3) to examine changes in AET and Deficit between past (c. 1700), present (1971–2000) and future (2020–49) climatological means derived from proxies, observations and projections; and (4) to infer how the magnitude of changing Deficit may contribute to changes in forest structure and composition. Location Yosemite National Park, California, USA. Methods We calculated the water balance within Yosemite National Park using a modified Thornthwaite‐type method and correlated AET and Deficit with tree species distribution. We used input data sets with different spatial resolutions parameterized for variation in latitude, precipitation, temperature, soil water‐holding capacity, slope and aspect. We used climate proxies and climate projections to model AET and Deficit for past and future climate. We compared the modelled future water balance in Yosemite with current species water‐balance ranges in North America. Results We calculated species climatic envelopes over broad ranges of environmental gradients – a range of 310 mm for soil water‐holding capacity, 48.3°C for mean monthly temperature (January minima to July maxima), and 918 mm yr^?1 for annual precipitation. Tree species means were differentiated by AET and Deficit, and at higher levels of Deficit, species means were increasingly differentiated. Modelled Deficit for all species increased by a mean of 5% between past (c. 1700) and present (1971–2000). Projected increases in Deficit between present and future (2020–49) were 23% across all plots. Main conclusions Modelled changes in Deficit between past, present and future climate scenarios suggest that recent past changes in forest structure and composition may accelerate in the future, with species responding individualistically to further declines in water availability. Declining water availability may disproportionately affect Pinus monticola and Tsuga mertensiana. Fine‐scale heterogeneity in soil water‐holding capacity, aspect and slope implies that plant water balance may vary considerably within the grid cells of kilometre‐scale climate models. Sub‐grid‐cell soil and topographical data can partially compensate for the lack of spatial heterogeneity in gridded climate data, potentially improving vegetation‐change projections in mountainous landscapes with heterogeneous topography.     

Modelling the influence of change in fire regime on the local distribution of a Mediterranean pyrophytic plant species (Cistus salviifolius) at its northern range limit

Aims To assess the potential distribution of an obligate seeder and active pyrophyte, Cistus salviifolius, a vulnerable species in the Swiss Red List; to derive scenarios by changing the fire return interval; and to discuss the results from a conservation perspective. A more general aim is to assess the impact of fire as a natural factor influencing the vegetation of the southern slopes of the Alps. Location Alps, southern Switzerland. Methods Presence–absence data to fit the model were obtained from the most recent field mapping of C. salviifolius. The quantitative environmental predictors used in this study include topographic, climatic and disturbance (fire) predictors. Models were fitted by logistic regression and evaluated by jackknife and bootstrap approaches. Changes in fire regime were simulated by increasing the time‐return interval of fire (simulating longer periods without fire). Two scenarios were considered: no fire in the past 15 years; or in the past 35 years. Results Rock cover, slope, topographic position, potential evapotranspiration and time elapsed since the last fire were selected in the final model. The Nagelkerke R² of the model for C. salviifolius was 0.57 and the Jackknife area under the curve evaluation was 0.89. The bootstrap evaluation revealed model robustness. By increasing the return interval of fire by either up to 15 years, or 35 years, the modelled C. salviifolius population declined by 30–40%, respectively. Main conclusions Although fire plays a significant role, topography and rock cover appear to be the most important predictors, suggesting that the distribution of C. salviifolius in the southern Swiss Alps is closely related to the availability of supposedly competition‐free sites, such as emerging bedrock, ridge locations or steep slopes. Fire is more likely to play a secondary role in allowing C. salviifolius to extend its occurrence temporarily, by increasing germination rates and reducing the competition from surrounding vegetation. To maintain a viable dormant seed bank for C. salviifolius, conservation managers should consider carrying out vegetation clearing and managing wild fire propagation to reduce competition and ensure sufficient recruitment for this species.     

Distribution patterns of boreal marshland birds: modelling the relationships to land cover and climate

Aim The aims of this work were (1) to study how well land‐cover and climatic data are capable of explaining distribution patterns of ten bird species breeding and/or feeding primarily on marshes and other wetlands and (2) to compare the differences between red‐listed and common marshland species in explanatory variables, and to study the predictability of their distribution patterns. Location Finland, northern Europe. Methods The data of the bird atlas survey carried out in 1986–89 using a 10 × 10 km uniform grid system in Finland were used in the analyses. Land‐cover data based on CORINE (Coordination of Information on the Environment) classification and climatic variables were compiled using the same 10 × 10 km grid. Generalized additive models (GAM) with a stepwise selection procedure were used to select relevant explanatory variables and to examine the complexity of the response shapes of the different species to each variable. The original data set was randomly divided into model training (70%) and model evaluation (30%) sets. The final models of common and red‐listed bird species richness were validated by fitting them to the model evaluation set, and the correlation between observed and predicted species richness was calculated. We assessed the discrimination ability of the binary models (single species) with the area under the curve (AUC) of a receiver operating characteristic (ROC) plot and the Kappa coefficient. Results Cover of marshland, shoreline length and mean temperature in April–June were significantly (P < 0.01) related to the common marshland species richness. Cover and clumping of marshland and mean temperature and precipitation in April–June were selected in the model of red‐listed marshland species richness. The level of discrimination in our single species models varied in ROC from fair to excellent (AUC values 0.70–0.95). Cover of marshland was included in all GAM models built for the target species, but clumping of marshland, shoreline length and cover of mires also appeared as important predictors in single species models. Seven species had statistically significant relationships with climatic variables in the multivariate GAMs. Cover of marshland was highest in squares in which the red‐listed bittern Botaurus stellaris, marsh harrier Circus aeruginosus and great reed warbler Acrocephalus arundinaceus and the water rail Rallus aquaticus were observed. Main conclusions Cover of marshland was the only variable which was included in all the models, reinforcing the close connection between the studied species and marshlands. Broad‐scale clumping of marshlands was important for the red‐listed species, probably due to the much lower population sizes of red‐listed species than those of common species. Land‐cover data produced in CORINE seems to be well suited for modelling the distribution patterns of marshland birds. Although climatic variables also strongly affect the studied marshland birds, habitat availability plays a crucial role in their occurrence. The distribution patterns of marshland birds at the scale of 10 × 10 km reflect the interplay between habitat availability and direct climatic variables.     

Co‐occurrence patterns of trees along macro‐climatic gradients and their potential influence on the present and future distribution of Fagus sylvatica L.

Aim During recent and future climate change, shifts in large‐scale species ranges are expected due to the hypothesized major role of climatic factors in regulating species distributions. The stress‐gradient hypothesis suggests that biotic interactions may act as major constraints on species distributions under more favourable growing conditions, while climatic constraints may dominate under unfavourable conditions. We tested this hypothesis for one focal tree species having three major competitors using broad‐scale environmental data. We evaluated the variation of species co‐occurrence patterns in climate space and estimated the influence of these patterns on the distribution of the focal species for current and projected future climates. Location Europe. Methods We used ICP Forest Level 1 data as well as climatic, topographic and edaphic variables. First, correlations between the relative abundance of European beech (Fagus sylvatica) and three major competitor species (Picea abies, Pinus sylvestris and Quercus robur) were analysed in environmental space, and then projected to geographic space. Second, a sensitivity analysis was performed using generalized additive models (GAM) to evaluate where and how much the predicted F. sylvatica distribution varied under current and future climates if potential competitor species were included or excluded. We evaluated if these areas coincide with current species co‐occurrence patterns. Results Correlation analyses supported the stress‐gradient hypothesis: towards favourable growing conditions of F. sylvatica, its abundance was strongly linked to the abundance of its competitors, while this link weakened towards unfavourable growing conditions, with stronger correlations in the south and at low elevations than in the north and at high elevations. The sensitivity analysis showed a potential spatial segregation of species with changing climate and a pronounced shift of zones where co‐occurrence patterns may play a major role. Main conclusions Our results demonstrate the importance of species co‐occurrence patterns for calibrating improved species distribution models for use in projections of climate effects. The correlation approach is able to localize European areas where inclusion of biotic predictors is effective. The climate‐induced spatial segregation of the major tree species could have ecological and economic consequences.     

Neophyte species richness at the landscape scale under urban sprawl and climate warming

Aim Land use and climate are two major components of global environmental change but our understanding of their simultaneous and interactive effects upon biodiversity is still limited. Here, we investigated the relationship between the species richness of neophytes, i.e. non‐native vascular plants introduced after 1500 AD, and environmental covariates to draw implications for future dynamics under land‐use and climate change. Location Switzerland, Central Europe. Methods The distribution of vascular plants was derived from a systematic national grid of 1 km² quadrates (n = 456; Swiss Biodiversity Monitoring programme) including 1761 species, 122 of which were neophytes. Generalized linear models (GLMs) were used to correlate neophyte species richness with environmental covariates. The impact of land‐use and climate change was thereafter evaluated by projections for the years 2020 and 2050 using scenarios of moderate and strong changes for climate warming (IPCC) and urban sprawl (NRP 54). Results Mean annual temperature and the amount of urban areas explained neophyte species richness best, with a high predictive power of the corresponding model (cross‐validated D² = 0.816). Climate warming had a stronger impact on the potential increase in the mean neophyte species richness (up to 191% increase by 2050) than ongoing urban sprawl (up to 10% increase) independently from variable interactions and model extrapolations to non‐analogue environments. Main conclusions In contrast to other vascular plants, the prediction of neophyte species richness at the landscape scale in Switzerland requires few variables only, and regions of highest species richness of the two groups do not coincide. The neophyte species richness is basically driven by climatic (temperature) conditions, and urban areas additionally modulate small‐scale differences upon this coarse‐scale pattern. According to the projections climate warming will contribute to the future increase in neophyte species richness much more than ongoing urbanization, but the gain in new neophyte species will be highest in urban regions.